A tlantic G eology 217

Palichnology of the Lower Devonian Wapske Formation, Perth-Andover- Mount Carleton region, northwestern New Brunswick, eastern Canada

Yaojun Han* and Ron K. Pickerill Department o f Geology, University o f New Brunswick, Fredericton, New Brunswick E3B 5A3, Canada

Date Received December 13, 1994 Date Accepted February 7, 1995

The Lower Devonian Wapske Formation of the Tobique Group in the Perth-Andover-Mount Carleton region of north­ western New Brunswick forms part of the tectonostratigraphic Tobique Zone. In this region, the formation is 4000 to 9000 m thick and is interpreted as deep-marine turbiditic in origin. Seven sedimentary facies are recognized. These are: conglomerate facies (Facies 1); massive and parallel-laminated sandstone facies (Facies 2); thin- to thick-bedded sandstone with minor mudstone interbeds and/or mud-capped facies (Facies 3); thin-bedded sandstone and mudstone facies (Facies 4); thin-bed- ded, graded, fine-grained sandstone facies (Facies 5); thin-bedded, fine-grained sandstone and silty mudstone facies (Facies 6); and thin- and medium-bedded siltstone and mudstone facies (Facies 7). These facies are grouped into five facies associa­ tions reflecting a deep-marine fan system consisting of distributary channels, depositional lobes, lobe or fan fringes, and interchannel areas associated with a basin plain. A relatively diverse and well-preserved ichnofauna, consisting of 23 ichnogenera (41 ichnospecies) and three vernacular ichnotaxa, is systematically described, these comprising Arthraria Billings, 1872; Bergaueria Prantl, 1945; Chondrites von Sternberg, 1833; Cochlichnus Hitchcock, 1858; Cosmorhaphe Fuchs, 1895; Cruziana d’Orbigny, 1842; Dendrotichnium Hantzschel, 1975; Didymaulichnus Young, 1972; Helminthoida Schafhautl, 1851; Helminthopsis Heer, 1877; Hormosiroidea Schaffer, 1928; Monomorphichnus Crimes, 1970; Neonereites Seilacher, 1960; Palaeophycus Hall, 1847; Paleodictyon Meneghini in Murchison, 1850; Phycodes Richter, 1850; Protopaleodictyon KsiqJkiewicz, 1970; Protovirgularia M ’Coy, 1850; Rusophycus Hall, 1852; Skolithos Haldeman, 1840; Taenidium Heer, 1877; Uchirites Macsotay, 1967; Umfolozia Savage, 1971; two track forms and indeterminate scratch markings. Their palaeoenvironmental distribution within the se­ quence correlates well with the interpreted submarine fan complex and compares favourably with other previously docu­ mented ichnofaunas from deep-marine fans. Collectively, the ichnofauna of the Wapske Formation is characteristic of Seilacher’s (1967) Nereites ichnofacies.

La formation de Wapske du Devonien inferieur du groupe de Tobique dans la region de Perth Andover/mont Carleton du nord-ouest du Nouveau-Brunswick fait partie de la zone tectonostratigraphique de Tobique. Dans cette region, la formation a 4 000 a 9 000 m d’epaisseur et elle est interpretee comme une formation d’origine turbiditique abyssale. On reconnatt sept facies sedimentaires, notamment : le conglomerat (facies 1); un facies de gres lamine massif et parallele (facies 2); un facies de gres en couches epaisses ou minces interstratifie de mudstone en quantite mineure ou couronne de boue (facies 3); un facies de mudstone et de gres en couches minces (facies 4); un facies de gres a grains fins, profile, en couches minces (facies 5); un facies de mudstone vaseux et de gres a grains fins, en couches minces (facies 6); et un facies de mudstone et de microgres en couches minces et moyennes. Ces facies sont groupes en cinq associations de facies correspondant a un systeme en eventail abyssal constitue de defluents, de lobes sedimentaires, de franges en lobe ou en eventail et de zones inter-canaux associees a une plaine synclinale. Une ichnofaune relativement diversifiee et bien preservee, constitute de 23 ichnogenres (41 ichnoespeces) et trois ichnotaxa courants, est decrite de fayon systematique. Ceux-ci comprennent 1’Arthraria Billings, 1872; le Bergaueria Prantl, 1945; les Chondrites von Sternberg, 1833; le Cochlichnus Hitchcock, 1858; le Cosmorhaphe Fuchs, 1895; le Cruziana d’Orbigny, 1842; le Dendrotichnium Hantzschel, 1975; le Didymaulichnus Young, 1972; 1’Helminthoida Schafhautl, 1851; 1’Helminthopsis Heer, 1877; VHormosiroidea Schaffer, 1928; Ic Monomorphichnus Crimes, 1970; le Neonereites Seilacher, 1960; le Palaeophycus Hall, 1847; le Palaeodictyon Meneghini dans Murchison, 1850; le Phycodes Richter, 1850; le Protopaleodictyon Ksi^zkiewicz, 1970; le Protovirgularia M’Coy, 1850; le Rusophycus Hall, 1852; le Skolithos Haldeman, 1840; le Taenidium Heer, 1877; 1’ Uchirites Macsotay, 1967; V Umfolozia Savage, 1971; deux formes de pistes et des marques de grattage. Leur repartition paleoenvironnementale a l’interieur de la sequence correspond bien avec le complexe en eventail sous-marin interprets et elle se compare favorablement avec les autres ichnofaunes des eventails abyssaux documentees auparavant. Collectivement, l’ichnofaune de la formation de Wapske est caracteristique de l’ichnofacies Nereites de Seilacher (1967). [Traduit par la redaction] Introduction tectonostratigraphic Tobique Zone (Fig. 1) (St. Peter, 1978a, The study area is located in northwestern New Brunswick, 1979; Pickerill, 1986, 1991; Wilson, 1990). In this area, the eastern Canada, from Perth-Andover to Mount Carleton, a dis­ Lower Devonian (Lochkovian-Pragian) Wapske Formation of tance of approximately 93 km, and is part of the the Tobique Group, the subject of this contribution, underlies approximately 205 km^ and has been estimated by St. Peter ♦Present address: Department of Geology, University of Alberta, Edmonton, (1978a,b, 1979) to be between 4000 and 9000 m in thickness. Alberta T6G 2E3, Canada

Atlantic Geology 30,217-245(1994) 0843-5561/94/030217-29S5.35/0 2 1 8 H an and P ickerill

Fig. 1. Generalized tectonostratigraphic map of New Brunswick, eastern Canada, showing location of the study area (modified from Pickerill, 1986; Fyffe and Fricker, 1987; Wilson, 1990). P-A = Perth-Andover, RB = Riley Brook; N = Nictau; MCP = Mount Carleton Park.

Surficial exposure, however, is restricted essentially to road­ below storm wave base in an outer shelf-upper slope deposi­ sides in the Riley Brook, Nictau, Mount Carleton Provincial tional environment. In this paper we briefly assess the sedi- Park and Perth-Andover districts (Fig. 1). The most common mentology of the Wapske Formation in more detail than in sedimentary lithofacies are grey, greenish grey or green silt- Pickerill (1986, 1991) and, in particular, describe systemati­ stones interbedded with varying proportions of grey, greenish cally its contained ichnotaxa. grey and brown, fine- to medium-grained sandstones and mud­ Ichnological research within the adjacent stones or shales. tectonostratigraphic Aroostook-Matapedia Zone has been con­ Several authors (e.g., St. Peter, 1978b, 1979, 1981, 1982; ducted by Pickerill (1980, 1981,1987) and Pickerill etal. (1987), Skinner, 1982; Irrinki and Crouse, 1986) have proposed a rela­ but essentially on strata of Late Ordovician and Early Silurian tively shallow marine, quiet water depositional setting for the age. The ichnology of the Lower Devonian Wapske Formation, Wapske Formation, based essentially on the evidence of its con­ until now, remains unstudied, except for a short ichnotaxonomic tained faunal communities (see also Boucot and Wilson, 1994). commentary by Pickerill (1991) and a series of more detailed However, an alternative scenario was envisaged by Pickerill ichnotaxonomic studies by Han and Pickerill (1994a,b, 1995). (1986, 1991) who, on sedimentological and taphonomic con­ Most strata of the Tobique Group in the study area lack body siderations, interpreted the formation as having been deposited fossils; however, many beds contain relatively well-preserved A tlantic G eology 219 and abundant ichnofossils that are considered useful for Faunal evidence, as recently reviewed in Boucot and Wilson palaeoenvironmental interpretation, particularly in view of the (1994), suggests an earliest Lochkovian to latest Pragian age fact that such previous interpretations of the sequence are at for the Wapske Formation. considerable variance. The purpose of this contribution is there­ Seven sedimentary lithofacies have been recognized in the fore to describe the ichnofaunas from this region and to com­ Wapske Formation in the present study. These lithofacies, indi­ ment on their interpreted palaeoenvironmental context. vidually differentiated on the basis of lithology, bed thickness, sandstone/mudstone ratios, bed geometry and contacts and in­ Stratigraphy and sedimentology ternal sedimentary structures, are summarized in Table 2. They are very similar to several of the seven submarine fan facies The Lower Devonian Tobique Group was originally pro­ first proposed and described by Mutti and Ricci Lucchi (1972) posed to comprise a basal Costigan Mountain Formation and a and subsequently refined by Walker and Mutti (1973) and Ricci conformably overlying Wapske Formation (St. Peter, 1978b) Lucchi (1975a,b). Associations of these sedimentary facies are (Table 1). The constituent lithotypes in these formations are interpreted as representing particular depositional environments similar, although the relative proportions differ, the Costigan present in a deep-water submarine fan system. These lithofacies, Mountain Formation being dominated by volcanic rocks and their associations and depositional subenvironments will be de­ the Wapske Formation by siliciclastic strata. Initially, St. Peter scribed separately (Han and Pickerill, in preparation) but are (1978b) believed that the contact between these formations was currently available in Han (1995). For brevity, however, the char­ probably isochronous; however, faunal evidence later indicated acteristics of these subenvironments, based on our interpreta­ that the Wapske Formation is locally intercalated and coeval tions of individual facies associations, can be summarized as with the Costigan Mountain Formation (St. Peter, 1979, 1981) follows. and thus that the contact should be regarded as diachronous. Mid-fan distributary channels comprise massively bedded,

Table 1. Summary of lithostratigraphy and previously interpreted depositional environments of the Tobique Group. Numbers refer to: 1, Wilson, 1990. 2, Skinner, 1982. 3, Irrinki and Crouse, 1986. 4,5,6 and 7, St. Peter, 1978b, 1979, 1981 and 1982. 8, Pickerill, 1986.

System Group Form ation Lithologies Depositonal Environments Devonian Gabbro and granite Sedimentary Unit: grey to green, quartzose or 1,3: shallow water marine. argillaceous siltstone and fine- to medium-grained 2,4,5,6,7: terrestrial and calcareous to non-calcareous, micaceous, shallow water marine. quartzose to arkosic, sandstone; minor grey to 8: outer shelf-upper slope Wapske green shale and slate; deep water marine.

Basaltic Unit: green-grey basalt; minor diabase sills and dykes, slate, greywacke and rhyolites;

Lower Tobique Rhyolitic Unit: pink and grey massive and lami­ Devonian nated rhyolite, minor basalt, slate and diabase. Unit 1: pink, red and grey rhyolite, quartz-feldspar 1,4,6: terrestrial to shallow porphyry flows, tuffs, and pyroclastic breccia. water marine.

Costigan Unit 2: grey shale, argillite and greywacke; minor Mountain diabase, rhyolite and basalt.

Unit 3: green-grey basalt flows and tuffs, minor volcanogenic sediments and conglomerate. Silurian Perham Chambord Bk. Grey to pink limestone; red-brown silty limestone. 1,8: shallow marine shelf. Hazeldean Grey to green siltstone and sandstone,minor shale. 1,8: shallow marine shelf. 220 H an and P ickerill

Table 2. Summary of lithofacies of the Wapske Formation and their essential characteristics.

Lithofacies 1 2 3 4 5 6 7

grey matrix- fine- to coarse- fine- to medium-grained thinly interbedded graded fine-grained grey fine- to very m udstones, silt-

Main lithologies s u p p o rte d grained sand- sandstones with minor fine-grained sand- sandstones or silt- fine-grained sand- stones with minor

conglomerates stones without thin mudstone interbeds, s to n e s Sc m u d - stone with rare stones, siltstones Sc thin sanstone

m u d s to n e s typically with mud-top s to n e s m u d sto n e s dark grey mudstones in te rb e d s

Bed thickness 7 0 -3 4 0 cm 2 -> 1 0 0 cm 2 -5 0 cm 1-2 0 c m 1-1 0 c m 0 .5 -5 c m 1 -2 0 c m Sand/mud ratio 1 :0 1:0 1 :0 .2 1 :0 .4 1 :0 .2 1:1 1:5 Bed geometry N/A ta b u la r tabular or lenticular ta b u la r ta b u la r tabular or lenticular ta b u la r Bed base N/A s h a rp sh a rp sharp or erosive sharp and planar s h a rp s h a rp Bouma units N o T a , T b Tae, abe, a-d Tabc, be, b-d, b-e Ta; rare Tb-e, c-e, de T d e , T c -c T d e , T e Sedimentary crude parallel- m a s s iv e o r parallel-, cross-laminated, parallel-, cross- graded, parallel-, massive, parallel- massive, parallel- structures la m in a te d p a ra lle l- scours, flute casts, mud laminated, flute cross-, cross climbing- la m in a te d L am in ated

la m in a te d c la s ts casts, mud clasts laminated, flute casts

Interpreted mass flow, esp. high-density turbidity currents low-density high-density low-density hemipelagic, pelagic depositional debris flows turbidity currents tu rb id ity g ra v ity Sc tra c tio n turbidity currents Sc low-density processes c u rre n ts flow s Sc hemipelagic turbidity currents matrix-supported conglomerates (Facies 1) that only rarely crop mon occurrence of thin- to medium-bedded, laterally continu­ out in the study area. Clasts are dispersed within a muddy or ous sandstones and mudstones suggest that the overall deposi­ silty matrix. tional environment was probably that of a mid or outer fan re­ Mid-fan depositional lobes consist of thin-bedded, fine­ gion of a deep-sea fan complex. grained, graded sandstones or coarse-grained siltstones (Facies 5), in association with thin- to medium-bedded fine-grained Palaeoenvironmental distribution sandstones interbedded with mudstones (Facies 4), medium­ OF TRACE FOSSILS grained mud-capped sandstones (Facies 3) and rare medium- to thick-bedded, medium-grained, massive and parallel-lami­ Ichnofaunas in the Wapske Formation are represented by nated sandstones (Facies 2). Facies 2 is generally restricted to horizontal fodinichnia, pascichnia, agrichnia and fewer upper parts of individual sequences in which the thin-bedded, repichnia, and were produced by unknown representatives of fine-grained sandstone beds are dominant. Thickening and both deposit- and suspension-feeders (Table 3). The ichnotaxa, coarsening upward cycles, high sandstone/mudstone ratios and comprising 23 ichnogenera, are collectively characteristic of laterally continuous beds are characteristic of this the deep-water Nereites ichnofacies of Seilacher (1967) (Fig. subenvironment. 2). The ichnofauna is relatively diverse and abundant with Outer fan lobes or fan fringes are dominated by thin-bed­ Helminthopsis and Palaeophycus representing most trace fos­ ded sandstones and mudstones (Facies 4) accompanied by mud­ sils. Graphoglyptids (notably Paleodictyon, Protopaleodictyon, stones and graded fine-grained sandstones. Beds within these Cosmorhaphe) and pascichnia (such as Helminthopsis, sequences exhibit a thinning and fining upward tendency. Helminthoida, Uchirites, Palaeophycus) dominate the pre-depo- Interchannel deposits consist of thin-bedded (with rare me­ sitional ichnocoenosis. In contrast, fodinichnia (especially Chon­ dium-bedded), fine-grained, graded sandstones (Facies 5). Amal­ drites), domichnia (e.g., Bergaueria, Skolithos) and repichnia gamation of sandstone beds is very common. Interbedded sand­ (such as Umfolozia, Protovirgularia, Cruziana) typify the post- stones and mudstones (Facies 4), mudstones (Facies 7), and depositional ichnocoenosis. Collectively, the overall ichnofaunal thinly interbedded sandstones and mudstones (Facies 6) are also assemblage of the Wapske Formation is comparable to several present. The sequences show a thinning or minor thickening of those recorded from other deep-marine fans (see McCann upward trend. and Pickerill, 1988; Miller, 1991, 1993; Crimes and Crossley, Basin plain-distal outer fan associations are characterized 1991; Crimes et al., 1992; Crimes and Fedonkin, 1994; and by laterally continuous thin-bedded mudstones and silty mud­ references therein). This is consistent with our sedimentologi- stones with thin, fine-grained sandstone interbeds (Facies 7); cal interpretations that suggest that deposition of the formation mud-capped sandstones (Facies 3) may be locally present. No occurred in a deep-marine environment. specific cyclical sequences occur. The most important parameters governing the distribution Within the proposed deep-water submarine fan system, the of deep-marine organisms are sediment texture, energy levels, precise location of the actual feeder channel(s) of the subma­ substrate consistency and stability, oxygen levels at or immedi­ rine fan(s) cannot be assessed. However, it is likely that the ately below the sediment-water interface, availability of food, source was located to the east or southeast of the present study sedimentation rates and frequency of event deposition. The area, as most conglomerates (Facies 1) are present there and palaeoecological controls over ichnofacies have been extensively palaeocurrent indicators such as flutes and cross-lamination sug­ discussed (e.g., Crimes, 1970a; Kern and Warme, 1974; Corbo, gest a west-northwest direction of turbidity current flow. The 1979; Frey and Seilacher, 1980; Pickerill et al., 1984; Bromley rare presence of coarse-grained channelized deposits, and com­ etal., 1984;Ekdale, 1985; Fillion and Pickerill, 1990;Uchman, A tlantic G eology 221

Table 3. Characteristics of ichnotaxa from the Wapske Formation.

Ichnotaxa D S F Feeding Ethology Toponomy N Pre-/Post-

Arthraria antiquata T Filter Fo CXH R ? Bergaueria hemispherica M T ?Filter Cu/Dm EN C Post- B. aff. p e r a ta M T ?Filter Cu/Dm EN R Post- Chondrites furcatus T Deposit Fo EN C Post- C. targionii T Deposit Fo EN R Post- C h o n d rite s isp. form A T Deposit Fo CXH R ? C h o n d rite s isp. form B T Deposit Fo CXH R Post- C-ochlichnus anguineus T Deposit Rp CXH R ? Cosmorhaphe fuchsi T Deposit Ps CXH R Pre- C. sin u o sa T Deposit Ps CXH C Pre- Cruziana problematica M T Deposit Rp CXH C Pre- Dendrotichnium haentzscheli T Deposit Fo CXH R ?

Didymaulichnus lyelli T Deposit Rp/Ps CXH R ? Hehninthoida cf. labyrinthica T Deposit Ps CXH R Pre- H. miocenica T Deposit Ps CXH C Pre- Helminthopsis abeli T M Deposit Ps CXH A Pre- H. cf. g r a n u la ta T M Deposit Ps CXH A Pre- H. hieroglyphica T M Deposit Ps CXH R Pre- Helminthopsis isp. T M Deposit Ps CXH A Pre- Hormosiroidea cf. beskidensis T ? Rp CXH C '/Pre- Monomorphichnus multilineatus T - Rp/Ps CXH R ‘/Post- Neonereites biserialis T Deposit Ps CXH C Post-

N. multiserialis T Deposit Ps CXH A Post- N. uniserialis T Deposit Ps CXH C Post-

Palaeophycus heberti T ?F ilter Ps CXH C Pre- P. striatus T ?Filter Ps CXH C Pre-

P. tubularis T ‘/Filter Ps CXH A Pre- Palaeophycus isp. T '/Filter Ps CXH A Pre- Paleodictyon isp. T M Deposit Ag CXH C Pre- Phycodes flabellus T Deposit Fo CXH C Post- P. palmatus T Deposit Fo CXH C Post- P. aff. p a lm a tu s T Deposit Fo CXH/EN R Post- P. p e d u m T Deposit Fo CXH C Post- P. tem p lu s T Deposit Fo CXH A Post- Protopaleodictyon incompositum T M Deposit Ag CXH/COH R Pre- Protovirgularia dichotoma T M Deposit Rp/Fo? CXH A '/Post- R u s o p h y c u s cf. carbonarius T M Deposit Cu CXH R Pre- Skolithos linearis M T Filter Dm EN A Post- Taenidium serpentinum M T Deposit Fo/Ps CXH R Post- Uchirites implexus T Deposit Rp/Ps CXH C Pre- U m fo lo zia cf. sin u o sa T ? Rp NA R Post-

Key: D = deep-water preference, S = shallow-water preference, F = facies crossing (T = typical, M = minor); Ethology: Ag = agrichnia, Cu = cubichnia, Dm = domichnia, Fo = fodinichnia, Ps = pascichnia, Rp = repichnia; Toponomy: CXH = convex hyporelief, COH = concave hyporelief, EN = endorelief, NA = not available; N = number of trace fossils (A = abundant (>10), C = common (3-10), R = rare (<3)); Pre- /Post- = pre - and post-depositional origin. 222 H an and P ickerill

Fig. 2. Schematic representation showing distribution of the common trace fossils within the deep-sea fan of the Wapske Formation (no scale implied). Numbers represent: 1. Arthraria, 2. Bergaueria, 3. Chondrites, 4. Coch/ichnus, 5. Cosmorhaphe, 6. Cruziana, 7. Didymaulichnus, 8. Helminthoida,9. Helminthopsis, 10. Hortnosiroidea, W.Monomorphichnus, \2. Neonereites, 13. Palaeophycus, 14. Paleodictyon, 15. Phycodes, 16. Protopaleodictyon, 17. Protovirgularia, 18. Rusophycus, 19. Skolilhos, 20. Taenidium, 21. Uchirites, 22. Track form B, 23. Scratch marks. Note: Track form A, Dendrotichnium haentzscheli and Umfolozia cf. sinuosa are not included as a result of their occurrence in small and isolated exposures that cannot be interpreted in terms of a specific palaeoenvironment (see also Table 4).

1992; Miller, 1993) and will not be repeated here. The are, in most examples, important with respect to toponomic pres­ palaeoenvironmental distribution of trace fossils in the Wapske ervation of trace fossils. The preservation of fine details in Formation has been directed toward the interpreted depositional Palaeophycus striatus, Phycodes palmatus and Rusophycus cf. environments as determined by sedimentological analysis (Table carbonarius indicates that the muds were relatively cohesive, 4) and is summarized below. as also suggested by the sharp burrow outlines of the associated The distributary channel facies association is devoid of body ichnotaxa (cf. Fillion and Pickerill, 1990). fossils, trace fossils and bioturbation. The primary reasons for The lobe or fan fringe facies association contains the few­ this are the coarse grain size and high-energy deposition, as est number of ichnotaxa (excepting the distributary channel fa­ well as the absence of shale interbeds precluding possible pres­ cies association). Helminthopsis and Palaeophycus are the only ervation of interface trace fossils. two types of traces found, being produced, respectively, by de­ The mid-fan depositional lobe facies association possesses posit- and suspension-feeders. Such a low diversity and abun­ a relatively diverse and abundant ichnocoenosis comprising dance may not, however, be representative of the original traces produced by both suspension- and deposit-feeders that ichnocoenoses, because lobe fringe deposits are not well repre­ were responsible for the formation of cubichnia, domichnia, sented in the sequence or, where present, are typically poorly fodinichnia, pascichnia and agrichnia. The most common forms exposed. Additionally, their component sandstones and/or silt- are horizontal agrichnia and pascichnia such as Cosmorhaphe, stones are commonly amalgamated, without shale interbeds, Helminthopsis, Palaeophycus, Paleodictyon and thereby suggesting that surface and shallow traces may have Protopaleodictyon, with fewer examples of cubichnia been eroded by turbidity currents. (Rusophycus), domichnia (Bergaueria) and fodinichnia The interchannel facies association exhibits the most di­ (Phycodes). The relative abundance of agrichnia and pascichnia verse and abundant ichnotaxa, comprising 18 ichnogenera and in the sequences is suggestive of a deep-water regime. The var­ one unnamed form (36 ichnospecies), including vertical ied ethological groupings and the existence of both deposit- and domichnia (Bergaueria and Skolithos), horizontal repichnia suspension-feeding ichnofaunas suggest the presence of suffi­ (Cochlichnus, Cruziana, Didymaulichnus, Hormosiroidea, cient nutrient levels and adequate oxygenation both within the Protovirgularia and scratch marks), fodinichnia (Chondrites, water column and at, or slightly below, the sediment-water in­ Phycodes) and pascichnia (Cosmorhaphe, Helminthoida, terface. Most trace fossils are preserved in thinly interbedded, Helminthopsis, Neonereites, Palaeophycus, Uchirites). The high fine-grained sandstone and mudstone lithofacies (Facies 4), diversity and abundance of deposit-feeding traces presumably whereas associated lithofacies only exhibit Palaeophycus and reflects the existence of an abundant nutrient supply and suffi­ Helminthopsis. Therefore, the sandstone and mudstone couplets cient oxygen levels to support a wide variety of infaunal and A tlantic G eology 223

Table 4. Palaeoenvironmental distribution of trace fossils from epifaunal benthos. The occurrence of suspension feeders the Wapske Formation. (domichnia) also suggests the existence of bottom currents car­ rying a continuous supply of suspended nutrients onto the sea floor. The preservation of surficial repichnia may reflect a re­ Environments Ichnotaxa duced erosive capacity of turbidity currents and/or less inten­ sive bioturbation by infaunal animals. Phycodes and the hori­ Basin plain- Helminthopsis abeli, H. isp., zontal repichnia display sharp, and only rarely lined bound­ distal outer fan Palaeophycus isp., Track form B aries, thereby indicating a relatively cohesive muddy substrate (cf. Fillion and Pickerill, 1990). Interchannel deposits are char­ Lobe fringes Helminthopsis isp., acterized by thin-bedded sandstones and mudstones (Facies 4) Palaeophycus isp. and thin-bedded, fine-grained, graded sandstones (Facies 5), which provide ideal preservational conditions for the traces. Bergaueria hemispherica, From an environmental aspect, interchannel areas are more Cosmorhaphe fuchsi, stable than channel, fan fringe and depositional lobe areas. In Helminthopsis aheli, addition to preservational considerations, this could perhaps Depositional H. hieroglyphica , H. isp., explain the presence of a presumably abundant and diverse epi- lobes Palaeophycus striatus, P. isp., fauna and infauna. Crimes and Fedonkin (1994) recently con­ cluded that interchannel areas associated with inner fans pos­ in mid-fan Paleodictyon isp., P h y c o d e s sessed diverse “shallow-water” trace fossils but fewer “deep­ templus, Protopaleodictyon water” ichnotaxa, whereas those in outer fans had diverse “deep­ incompositum, Rusophycus cf. water” trace fossils and a general absence of “shallow-water” carbonarius trace fossils. Interchannel deposit sequences from the Wapske Formation exhibit mostly facies-crossing forms and more or Arthraria antiquata, Bergaueria less equal representatives of typical “shallow” and “deep” wa­ hemispherica, B. aff. p e ra ta , ter forms. Chondrites furcatus, The basin plain-distal outer fan facies association comprises essentially homogeneous muddy strata with sporadic turbidite C. targionii, C. isp. form A and sandstone interbeds. Exposed sequences reveal only isolated B, Cochlichnus anguineus, examples of Helminthopsis, Palaeophycus and unassigned Cosmorhaphe fuchsi, C. sinuosa, tracks. The scarcity of ichnotaxa may reflect incomplete pres­ Cruziana problematica, ervation of trace fossils because of the lack of turbidite sand­ Didymaulichnus lyelli, stones as a casting medium, or may be a result of the environ­ ment being inimical to the support of a benthic population ei­ Helminthoida cf. labyrinthica, ther as a result of low nutrient levels or depressed oxygen sup­ H. miocenica, Helminthopsis plies at, or within, the upper levels of the surficial substrates. a b eli, H . cf. g ra n u la ta , In a general context, thin-bedded sandstones and siltstones H. hieroglyphica, H. isp., contain considerably more ichnotaxa, both in terms of diversity Interchannel Hormosiroidea cf. beskidensis, and abundance, than do the thick-bedded turbidites, similar to areas Monomorphichnus analogous Cretaceous turbidites of northern California described by Miller (1993). The submarine fan system of the Wapske For­ multilineatus, Neonereites mation appears to have contained sufficient food supplies and biserialis, N. multiserialis, oxygen levels to support a widespread population of benthic N. uniserialis, Palaeophycus organisms. However, oxygen levels in sediment below the sedi­ heberti, P. striatus, P. tubularis, ment-water interface were at times presumably sufficiently re­ P. isp., Phycodes flabellus, duced so that only the producer of Chondrites occurred in the P. palmatus, P. aff. p a lm a tu s, sequence (see also Bromley and Ekdale, 1984; Hakes, 1985; Savrda and Bottjer, 1986). Secondary factors, such as mode of P. pedum, P. templus, preservation and degree of exposure and weathering, also po­ Protovirgularia dichotoma, tentially influence the observed ichnocoenoses. In newly ex­ Skolithos linearis, Taenidium posed outcrops where both talus and weathering processes are serpentinum, Uchirites implexus, relatively minor, detailed examination and sampling of trace scratch marks fossils was more difficult compared to outcrops with extensively exposed and weathered talus. For example, distributary chan­ Note: Dendrotichnium haentzscheli, Umfolozia cf. sinuosa and Track nel deposits near Nictau Lake and depositional lobe deposits form A are not included as a result of their occurrence in small and near Perth-Andover revealed no, or only sporadic, trace fossils isolated exposures that cannot be interpreted in terms of a specific because of the generally poor exposure and smaller outcrop size, palaeoenvironment (see also Fig. 2). resulting in a deficiency of talus. Secondly, traces are preserved 224 H an and P ickerill toponomically as semireliefs and, for optimum preservation, a tions. For completeness, however, each of these ichnotaxa are contrast in lithology between the excavated and casting sedi­ figured herein. Ichnotaxa described in open nomenclature are ments is required (Fillion and Pickerill, 1990). In the Wapske described separately. Formation, both monolithologic sandstones (lobe deposits and The preservational styles and essential characteristics of portions of lobe fringe deposits) and shales (basin plain depos­ each form are listed in Table 3. Table 4 summarizes the its) display the fewest number of trace fossils and a realistic palaeoenvironmental distribution in the sequence. Specimens scenario of the ichnocoenoses is difficult to reconstruct. How­ with prefix TF and TF.F are housed in the Department of Geol­ ever, couplets of thinly interbedded sandstones and mudstones ogy, University of New Brunswick, Fredericton, New Brunswick, provide an ideal medium for the preservation of trace fossils Canada; the prefix NBMG represents specimens deposited in and the degree of weathering and talus development in these the Division of Natural Sciences, New Brunswick Museum, Saint rock types provide ideal conditions for observations of semirelief John, New Brunswick, Canada. traces, such as with the lobe fringe and interchannel deposits. Ichnogenus Arthraria Billings, 1872 C onclusions Type ichnospecies: Arthraria antiquata Billings, 1872 The main conclusions that can be drawn from this study Arthraria antiquata Billings, 1872 are: Figure 3A (1) The Wapske Formation of the Lower Devonian (Lochkovian-Pragian) Tobique Group, located between Perth- Material: Four specimens (TF9208-23, TF9208-30, TF9210- Andover and Mount Carleton, northwestern New Brunswick, 10, TF112). possesses seven sedimentary lithofacies that can be organized into five facies associations. These facies associations reflect Description: Dumb-bell shaped traces preserved in convex deposition in distributary channels, depositional lobes, lobe or hyporelief on 26 to 30 mm thick, grey, parallel-laminated fine­ fan fringes, interchannel and basin plain subenvironments in grained sandstone or siltstone. Each consists of a shallow and association with a deep-water submarine fan complex. narrow stem connecting two wider and more deeply impressed (2) A relatively diverse and well-preserved ichnofauna is, terminations which are heart-, kidney-, or arrowhead-shaped. to varying degrees, present in these various subenvironments. The stems are about 3 to 30 mm long and 2 to 11 mm wide and In total these ichnofaunas include 23 ichnogenera (41 terminations 4 to 11 mm long and 2 to 11 mm wide. Termina­ ichnospecies) and three vernacular forms. Their tions do not extend upward to form U-shaped tubes and the palaeoenvironmental distribution, as determined from facies burrows possess no spreiten. analysis, suggests close comparisons with other ichnofaunas described from deep-water submarine fan systems. The Remarks: Fillion and Pickerill (1984) reviewed and redefined ichnofauna of the Wapske Formation is, collectively, character­ the monospecific ichnotaxon Arthraria, distinguishing it from istic of Seilacher’s (1967) Nereites ichnofacies. The main pri­ the morphologically similar, at least in basal expression, mary controlling factors of the ichnofaunal distribution were Bifungites Desio, which, however, has vertical tubes without a food supply, oxygen levels, sediment texture, sedimentation rates spreite, and Diplocraterion Torell which has both vertical tubes and frequency of event deposition. and a spreite. Arthraria is believed to have been produced by a (3) Secondary factors considered important with respect to worm or wormlike organism (Fillion and Pickerill, 1990). Ter­ trace fossil distributions in the various subenvironments of the minations of different shapes may occur in this ichnospecies, Wapske Formation include nature and quality of outcrop, such even within a single specimen. Size of individual specimens as weathering characteristics, and amount of talus available for may also vary considerably. Thus all reliable specimens of detailed analysis. Arthraria should be included within Arthararia antiquata (Fillion and Pickerill, 1984). Systematic ichnology Ichnogenus Bergaueria Prantl, 1945 In accordance with common ichnological procedure (Hantzschel, 1975; Pemberton and Frey, 1984; Fillion and Type ichnospecies: Bergaueriaperata Prantl, 1945 Pickerill, 1990), the ichnotaxa in this contribution are described in alphabetical order rather than in any morphological or The ichnotaxonomy of single-entrance, plug-shaped, soft- behavioural classification groupings (e.g., as in Osgood, 1970; substrate trace fossils has been reviewed in detail by Pemberton Ksiqzkiewicz, 1977). This procedure is adopted for ease of ref­ et al. (1988). By its much larger diameter/height ratio, erence to a particular ichnotaxon and also because in practice Bergaueria can be distinguished from Conichnus Myannil, many ichnotaxa may be indicative of more than one behavioural Dolopichnus Alpert and Moore and Conostichus Lesquereux activity. For those ichnotaxa we have previously considered in by its lack of a distinct apical disc and wall ornamentation detail (Han and Pickerill, 1994a,b, 1995), namely Helminthopsis (Pemberton etal., 1988). ispp., Phycodes templus and Protovirgularia dichotoma, we only Based mainly on characteristics of the distal termination include our previous diagnoses and avoid repetitious descrip­ and the presence or absence of wall linings, Pemberton et al. A tlantic G eology 225

Fig. 3. Trace fossils from the Wapske Formation. (A) Arthraria anliquala (TF9208-23) (arrowed), x 1.14. (B) Bergaueria hemispherica (TF9209- 20), x 0.77. (C, D) Bergaueria aff. perata (TF 9201-21), x 1.0 (C, top view; D, lateral view). (E) Bergaueria hemispherica (TF9208-24), x 0.86 (vertical section showing the V-shaped laminae). (F) Cochlichnus anguineus (TF.F118) (arrowed) intergradational with Helminthopsis isp., x 1.38. (G) Chondrites furcalus (TF.F223), x 0.83. (H) Chondrites targionii (TF9307-04), x 0.82. A-F are preserved in convex hyporelief; G and H in endorelief.

(1988) recognized four ichnospecies of Bergaueria, namely, B. of these have been discussed by Pemberton et al. (1988) and perata, B. langi (Hallam), B. radiata Alpert and .8. hemispherica Pickerill (1989). Crimes, Legg, Marcos and Arboleya. Ethological considerations 2 26 H an and P ickerill

Bergaueria hemispherica Crimes, Legg, Marcos and miform worms (Osgood, 1970; Howard and Frey, 1984), Arboleya, 1977 arthropods (Ekdale, 1977), anthoptiloid sea pens (Bradley, 1981) Figure 3B,E and nematodes (Bromley and Ekdale, 1984; Howard and Frey, 1984; Swinbanks and Shirayama, 1984). Material: Five specimens (TF9208-24, TF9208-1-13, TF9209- Ichnospecific classification of Chondrites is traditionally 20, TF.F172, TF.F58). based on width of the branches and mode and angle of branch­ ing (Ksiqzkiewicz, 1977), and has resulted in the description of Description: Vertical, cylindrical to dome-like burrows pre­ more than 170 ichnospecies (Chamberlain, 1971), many of served in convex hyporelief on the soles of 8 to 75 mm thick, which are extremely difficult, if not impossible, to differentiate grey, parallel- and cross-laminated siltstone and fine-grained or may, in fact, be simply ontogenetic variants (Hakes, 1976; sandstone. The burrows have steeply to gently inclined, Ksiqzkiewicz, 1977). unomamented outer margins. The basal surfaces are flat or Typically occurring infaunally beneath an oxygen-deficient rounded without typical depressions or ridges. Dimensions are sea-floor below the sediment-water interface (Bromley and variable. Diameter is equal to or generally greater than height. Ekdale, 1984), Chondrites has commonly been regarded as an The distal ends are 15 to 28 mm in diameter, the proximal ends indicator of dysaerobic or even anoxic conditions (Bandel, 1973 ; 18 to 35 mm, and the heights 8 to 12 mm; the average ratio of Bromley and Ekdale, 1984, 1986). The presence of Chondrites diameter/height is between 2.5-1. The burrow-fill is the same alone in a sedimentary unit, its burrow diameter and the degree as the host rock and no lining is observed in vertical section. of branching can all be used to interpret changes in environ­ However, in one vertically sectioned specimen, the primary par­ mental parameters (Hakes, 1985; Savrda and Bottjer, 1986). allel laminae are distorted to a V-shape pointing to the burrow base (Fig. 3E), which has not been described previously. Chondrites furcatus von Sternberg, 1833 Figure 3G Remarks: Bergaueria hemispherica differs from B. perata and B. radiata by the absence of a central depression and radial Material: Three specimens (TF9211-12, TF.F223, TF.F227). ridges and from B. langi by the absence of a thick burrow wall (Pemberton etal., 1988). Description: The burrow systems are well-preserved in endorelief within greenish grey, fine-grained, 10 to 15 mm thick, Bergaueria aff. perata Prantl, 1945 sandstones. The burrows are dendritic, maximum width 95 mm, Figure 3C,D ramifying symmetrically or asymmetrically at very acute angles, 25° to 55°, commonly 30° to 40°. Mostly they bifurcate, but Material: One specimen (TF9201-21). occasionally trifurcate. There are up to 3 or 4 orders of branch­ ing. The interval length of branching is irregular, either long Description: Hemispherical burrow preserved in convex or short. At branching sites no expansion is present. Individual hyporelief on the sole of a 29 mm thick siltstone. The burrow branches are cylindrical, horizontal or inclined, straight or has a smooth, unornamented, gently inclined outer wall and a curved, but never bend, wind, cross or interpenetrate. Burrow flat apical base which has a circular depression. In plan view, fill is lighter in colour and finer in grain size than the host the specimen is circular with an upper diameter of 26 mm, a rock. The burrow-fill is separated into transverse segments, lower diameter of 8.5 mm, and a height of 12 mm. The base of which are individually 2.5 to 5 mm (commonly 4 mm) long the specimen is flat, symmetric and possesses a small depres­ and 2 to 2.5 mm wide. The boundaries of the transverse seg­ sion, 3 to 3.5 mm in width. The burrow-fill is massive and of ments are either straight or meniscate without unique form. the same grain size as the host rock. A thin lining is present. There is no change in width along the branches.

Remarks: Pickerill (1989) provided an in-depth discussion on Remarks: The present specimens are very similar to Chondrites this ichnospecies. The present specimen has a flat base with a type B of Osgood (1970) and virtually identical to C. furcatus central depression, but no radial ridges as in B. perata to which, of Ksiqzkiewicz (1977) and Crimes and Crossley (1991), and, however, it is otherwise most comparable. The central depres­ therefore, are considered conspecific. Chamberlain (1977) re­ sion differentiates it from B. hemispherica. garded C. furcatus as a junior synonym of C. recurvus Brongniart. However, C. furcatus differs from C. recurvus by Ichnogcnus Chondrites von Sternberg, 1833 the former’s more asymmetrical dichotomous branching pat­ tern and the latter’s more symmetrical dichotomous branches Type ichnospecies: Fucoides antiquus Brongniart, 1828 which are also more regular and outward curving. Addition­ ally, the present specimens do not exhibit the prominent Chondrites has previously been interpreted as both a feed­ recurvature characteristic of C. recurvus. C. furcatus differs from ing and dwelling burrow and probably indicates a systematic C. targionii (Brongniart) by its curved branches and narrower search for food by repeated probing of sediment (Vossler and branching angles and from C. intricatus (Brongniart) by its Pemberton, 1988). Previously proposed tracemakers include a larger size and predominance of asymmetrical dichotomous variety of endobenthic deposit feeders such as sipunculids branching. (Simpson, 1957), multitentacled organisms (Taylor, 1967), ver­ A tlantic G eology 227

Chondrites targionii (Brongniart, 1828) Remarks: The poorly preserved and incomplete specimen was Figure 3H found only by splitting the sandstone. Its distinctive fan-like branching system with slender branches distinguishes it from Material: Two specimens (TF9307-04, TF115). the other ichnospecies of Chondrites described here.

Description: The specimens are well-preserved in endorelief Ichnogenus Cochlichnus Hitchcock, 1858 in a greater than 50 mm thick, grey, parallel-laminated, fine­ grained sandstone. A maximum of five orders of branching are Type ichnospecies: Cochlichnus anguineus Hitchcock, 1858 present, resulting in a straight-branched dendritic pattern up to Cochlichnus anguineus Hitchcock, 1858 127 mm across. The burrow systems are regular, asymmetric Figure 3F and loosely bifurcated. Branches regularly alternately diverge from a “main” stem with branching angles ranging from 30° to Material: One specimen (TF.F118). 50°. Branches are straight, 2 to 7 mm wide, constant or taper­ ing and 7 to 53 mm long. Burrow fill is black mudstone which Description: Smooth, unbranched sine-like trail preserved in is finer in grain size and darker in colour than the host matrix. convex hyporelief on the sole of a 25 mm thick, brown, fine­ grained sandstone. The trail, 16 mm long and 1.5 mm wide, Remarks: The present specimens are regarded as Chondrites exhibits a wavelength of 2 mm and amplitude of 4 mm. targionii, because they show a dendritic branching pattern, strong or slender, straight, uniformly wide or slightly tapering Remarks: Cochlichnus is usually regarded as a repichnion branches, and angular branching patterns. It differs from C. (Hitchcock, 1858; Pienkowski and Westwalewicz-Mogilska, furcatus, C. intricatus and C. recurvus by its straighter branches, 1986), but true burrows have also been documented (Webby, and from C. intricatus by its larger size, smaller branching 1970; Eagar et al., 1985). It is thought to have been produced angles, and possession of asymmetrical dichotomous branches. by nematodes and annelids (Hitchcock, 1858; Seilacher, 1963; It also differs from C. succulens which is characterized by ir­ Moussa, 1969,1970; Hakes, 1976). Elliott (1985) demonstrated regular, nodose, curved to tortuous burrows. the geometric nature of the repeated clothoid curves of Cochlichnus kochi Ludwig, concluding that this enabled effi­ Chondrites isp. form A cient propulsion through watery surface muds of an animal with Figure 4B a hydrostatic skeleton. The present specimen is directly connected to Helminthopsis Material: One specimen (TF9208-43). isp. as a compound specimen (Pickerill, 1994). The regular sine curve changes to an irregularly winding structure presumably Description: The tiny, tree-like specimen is poorly preserved as a result of change in the tracemaker’s behaviour. in convex hyporelief on the surface of a 75 mm thick, grey, parallel-laminated, fine-grained sandstone. It consists of a very Ichnogenus Cosmorhaphe Fuchs, 1895 small, horizontal to inclined, symmetrically bifurcated branch­ ing system, up to 20 mm across and possessing two orders of Type ichnospecies: Cosmorhaphe sinuosa Azpeitia Moros, branching. Branching sites have no swelling and are subangular. 1933 Branching angles vary between 30° to 75°. Burrows are slen­ Cosmorhaphe fuchsi Ksiazkiewicz, 1970 der, straight to slightly wavy, about 1 mm wide and 3 to 6 mm Figure 4C long. Branch width is uniform throughout the whole burrow system. Material: Three specimens (TF9202-16, TF9208-41, TF9208- 1-30). Remarks: The poor and incomplete preservation of the speci­ men precludes ichnospecific assignment. Description: Smooth, unbranched, composite meanders pre­ served in convex hyporelief on the soles of 40 to 85 mm thick, Chondrites isp. form B grey, massive to parallel-laminated, fine-grained sandstones. Figure 4 A The meanders of the first-order are high but wide at the base (wavelength 120 to 250 mm, amplitude 45 to 170 mm, wave- Material: One specimen (TF9211-10). length/amplitude ratio 1.47-2.67). Meanders of the second- order are very variable, deep or shallow, usually not compressed Description: Fan-shaped burrow system preserved in endorelief or constricted, regular to irregular (wavelength 25 to 80 mm, within a 20 mm thick, brown, fine-grained sandstone. It con­ amplitude 5 to 35 mm, wavelength/amplitude ratio changeable, sists of a slender, widely spaced, symmetrically dichotomous most at least larger than 2.5). The structures are continuous, branching system, 31 mm across. The almost straight branches each with almost a constant diameter of between 1 to 15 mm. are 0.5 to 1.2 mm wide and 20 to 50 mm long, with constant width along individual segments. A maximum of two orders of Remarks: The absence of compression and constriction of both branching are present. Branching junctions are without swell­ the first- and second-order meanders characterizes Cosmorhaphe ings and diverge with angles varying from 20° to 30°. Burrow- fuchsi. There are distinct difference between C. fuchsi, whose fill is dark grey mudstone. first- and second-order meanders are not compressed, and C. 2 2 8 H an and P ickerill

Fig. 4. Trace fossils from the Wapske Formation. (A) Chondrites isp. form B (TF9211-10), x 1.12. (B) Chondrites isp. form A (TF9208-43), x 1.0. (C) Cosmorhaphe fuchsi (TF9208-1-30), x 0.54. (D) Cruziana problematica (TF9209-18), x 1.93. (E) Dendrotichnium haentzscheli (TF9210- 2-11), x 1.0. (F) Didymaulichnus lyelli (TF9201-13), x 1.12. (G) Cosmorhaphe sinuosa (TF.F142), x 0.74. A is preserved in endorelief; the remainder in convex hyporelief. sinuosa (Azpeitia Moros), whose first-order meanders are com­ could live both on the sea-bottom or close to it, as indicated by pressed, but whose second-order meanders may or may not be present day Cosmorhaphe in the deep sea where it is interpreted restricted. Cosmorhaphe is a distinctive and typically deep-wa­ as an infaunal burrow system positioned just below the sedi­ ter flysch form that may be pre- or post-depositional in origin ment surface (Ekdale and Berger, 1978; Ekdale, 1980a, b). It (Ksiazkiewicz, 1977). It may be produced by animals which has been interpreted as a grazing trail (Webby, 1969; Hantzschel, A tlantic G eology 229

1975) or a feeding burrow (Seilacher, 1962). Any worm-shaped Cruziana (cf. Romano and Whyte, 1987; Pickerill and Peel, animal or gastropod without external ornamentation could be 1990; Pickerill, 1994; Keighley and Pickerill, 1995a). responsible for this trace (Ksiqzkiewicz, 1977). Ichnogenus Dendrotichnium Hantzschel, 1975 Cosmorhaphe sinuosa (Azpeitia Moros, 1933) Figure 4G Type ichnospecies: Dendrotichnium llarenai (Farres, 1967) Dendrotichnium haentzscheli (Farres, 1967) M aterial: One specimen (TF.F142). Figure 4E

Description: Smooth, unbranched, composite meanders, 1.8 mm Material: One specimen (TF9210-2-11). in diameter, preserved in convex hyporelief on the sole of a 25 mm thick, parallel-laminated, fine-grained sandstone. The first- Description: The specimen comprises a smooth, horizontal bur­ order meanders are narrow and high, 15 to 53 mm in wave­ row system preserved in convex hyporelief on the sole of a grey length and 130 mm in amplitude. These are superimposed by siltstone. The burrow consists of a central main stem and nearly second-order meanders which range from 25 to 37 mm in wave­ alternately branched lateral appendages. The burrow system length and 5 to 15 mm in amplitude. The second-order mean­ covers a width of 32 mm. The straight main stem is 4 mm wide ders vary in size within the specimen. and 63 mm long; the lateral appendages, 3 to 4 mm wide and 12 to 20 mm long, branch out obliquely at angles of 30° to 45° Remarks: Seilacher (1977) suggested that Cosmorhaphe and are constant in width. The burrow-fill is similar in grain sinuosa can be distinguished from other related forms by the size to the host rock. low amplitude of the secondary meanders and the relatively small burrow diameter. Cosmorhaphe sinuosa is related to C. Remarks: In contrast to truly dendritic burrow systems such as helminthopsoida differing only in the latter’s less regularly me­ Chondrites, the side branches of Dendrotichnium may alter­ andering course (Ksi^zkiewicz, 1977). nate regularly and never show any secondary branches (Seilacher, 1977). Three ichnospecies of Dendrotichnium have Ichnogenus Cruziana d’Orbigny, 1842 been described, namely D. haentzscheli, D. llarenai and D. alternans Seilacher. They are differentiated on the basis of ap­ Type ichnospecies: Cruziana rugosa d’Orbigny, 1842 pendage branching angles and patterns and nature of the cen­ Cruzianaproblematica (Schindewolf, 1921) tral stem. Dendrotichnium haentzscheli differs from/), llarenai Figure 4D by the latter’s right angled branched appendages and from D. alternans by the latter’s regularly alternately branched lateral Material: Nine specimens (TF9201-22, TF9209-18, TF.F162, appendages and zigzag meandering main stem. Dendrotichnium TF.F155). occurs most commonly in flysch deposits (Seilacher, 1977; Crimes etal., 1981). Description: Specimens are moderately or poorly preserved in convex hyporelief on the soles of 20 to 40 mm thick, brown or Ichnogenus Didymaulichnus Young, 1972 grey, parallel-laminated, fine-grained sandstones. They are straight or slightly curved, 8 to 26 mm long and 4 to 8 mm Type ichnospecies: Fraena lyelli (Rouault, 1850) wide, and composed of two lobes (symmetrical or slightly asym­ Didymaulichnus lyelli (Rouault, 1850) metrical), each 1.8 to 3,5 mm wide and 0.1 to 1.5 mm deep, Figure 4F separated by a very shallow median furrow more or less extend­ ing the length of the traces. The lobes are characterized by faint Material: One specimen (TF9201-13). or strong, closely spaced transverse or slightly oblique scratch marks which are neither bifid nor bunch together. Description: Narrow and elongate, curved, horizontal bilobate convex hyporelief, 7 to 8 mm wide and 53 mm long, preserved Remarks: Lessertisseur (1955) and Osgood (1970) reviewed on the sole of a maroon siltstone. A distinct median furrow, 0.5 the origin of Cruziana. It is believed that most were produced to 1 mm wide, runs the whole length. The symmetrical lobes by trilobites (Seilacher, 1970) though some may have been pro­ have no surface ornament. The specimen undulates slightly in duced by trilobite-like arthropods, notostracan branchiopods the horizontal plane and therefore the lobe margins are not (Bromley and Asgaard, 1979), aglaspidids (Fisher, 1978) or even strictly straight but wavy though still clearly defined. The mar­ vertebrates (Shone, 1978, 1979). Typically a shallow-marine gins of the lobes are steep, without marginal ridges. form, Cruziana has also previously been reported from fresh­ water (Bromley and Asgaard, 1979) and deep-marine (Pickerill Remarks: The smooth lobes in Didymaulichnus differentiates et al., 1987) deposits. Cruziana problematica differs from it from Cruziana which has scratch marks (Young, 1972). It is Didymaulichnus Young by the presence of scratch marks on commonly attributed to gastropods (Glaessner, 1969; Hakes, each lobe. It is morphologically identical to the ribbon forms of 1976) or trilobites (Crimes, 1970b; Bradshaw, 1981) and is a Isopodichnus Bornemann, though following Bromley and facies-crossing form (Young, 1972; Hakes, 1976; Eagar et al., Asgaard (1979), such forms should be regarded as synonyms of 1985). As the present specimen is characterized by an absence 230 H an and P ickerill

of oblique scratches, lateral ridges, marginal bevels or alternat­ other without a bow-like form or tendency to coil. Most mean­ ing step-wise compressed sections, it is assigned to ders bulge at their turning sites resulting in a horseshoe-like Didymaulichnus lyelli. form. The width of the meanders is between 1 to 2 mm, this remaining constant in individual specimens. The surface of the Ichnogenus Helminthoida Schafhautl, 1851 meanders is smooth.

Type ichnospecies: Helminthoida labyrinthica Heer, 1865 Remarks: These specimens are veiy similar to descriptions, and criteria for the recognition, of H. miocenica by Ksiqzkiewicz Seilacher (1977) erected the ichnogenus Helminthorhaphe (1977) and Crimes and Crossley (1991). The meanders of H. to include Helminthoida crassa Schafhautl, H. japonica Tanaka miocenica are less regular and tighter than in H. crassa. Simi­ and a new ichnospecies H. reflecta. However, Crimes et al., lar bulging meanders can also be observed in Ksiqzkiewicz’s (1981), Crimes and Anderson (1985) and Crimes and Crossley (1977, p. 166, figs. 36a,d,e) specimens. (1991) argued for retention of Helminthoida, their view being accepted, amongst others, by Benton (1982a), McCann and Ichnogenus Helminthopsis Heer, 1877 Pickerill (1988), Narbonne and Aitken (1990) as well as herein. Helminthoida is still, however, an ichnogenus that exhibits con­ Type ichnospecies: Helminthopsis magna Heer, 1877 siderable diversity in form and spacing of its meanders Helminthopsis abeli Ksiaikiewicz, 1977 (Ksiqzkiewicz, 1977) and course, and there is considerable con­ Figure 5C fusion regarding differentiation of several of its ichnospecies. Recently, Crimes and Crossley (1991) provided a detailed study M aterial: Seven specimens (NBMG 9670 - NBMG 9675, and of Helminthoida and placed H. japonica in synonymy with H. NBMG 9676 (partim)) and more than 50 specimens collected crassa. They also suggested that more or less regular and closely and many more field occurrences. spaced parallel meanders arranged in a bow-like forms should be included within H. crassa, while lower, less compressed me­ Emended diagnosis: Helminthopsis that is loosely winding or anders and a totally unbowed form with no spiral development meandering. Meanders irregular and variable in shape, mostly should be regarded as H. miocenica Sacco. For those with a deep, but also shallow, with bell-shaped and, or, horseshoe­ distinct tendency to coil they suggested accommodation within shaped segments, but no straight segments or loops. The axes H. labyrinthica. In our experience, this is a workable scheme of the meanders are not parallel. Diameter is variable and gen­ and is therefore adopted herein. Helminthoida is typically, erally constant within a single specimen (modified from though not exclusively, a deep-water trace fossil. Ksiqzkiewicz, 1977).

Helminthoida cf. labyrinthica Heer, 1865 Description and Remarks: See Han and Pickerill, 1995. Figure 5B Helminthopsis cf. granulata Ksiaikiewicz, 1968 Material: One specimen (TF.F178). Figure 5D

Description: Burrow is preserved in convex hyporelief on the Material: One specimen (NBMG 9682). sole of a 23 mm thick, buff, fine-grained sandstone. The mean­ ders are closely spaced and even partially overlap. The mean­ Emended diagnosis: Helminthopsis which is variably and ir­ ders vary in width between 2 to 9 mm (commonly 6 mm); me­ regularly but loosely meandering. Surface is characteristically ander height is 35 to 50 mm and intervals between them 0 to 4 covered with warts and ridges arranged parallel to its axis (modi­ mm. The burrow surface is smooth and burrow-fill is similar in fied from Ksiqzkiewicz, 1977). grain size to the enclosing sandstone. Description and Remarks: See Han and Pickerill, 1995. Remarks: The present specimen is only partially complete but is tentatively assigned to H. cf. labyrinthica by the presence of Helminthopsis hieroglyphica H eer in M aillard, 1887 tight meanders and its tendency to coil. Figure 5F

Helminthoida miocenica Sacco, 1886 Material: Six specimens (NBMG 9676 (partim), NBMG 9677 Figure 5A - NBMG 9681) and more than 44 specimens collected and many more field occurrences. Material: Three specimens (TF.F111, TF.F113, TF.F114). Diagnosis: Helminthopsis in which the windings, normally wide Description: Specimens are preserved in convex hyporelief and low, comprise straight segments interspersed with irregu­ on the soles of 20 to 27 mm thick, brown, fine-grained sand­ larly sinuous and variably developed segments. The tortuous stones. The meanders are up to 34 mm high while the distance segments may be bell-shaped but not horseshoe-shaped. The between the arms of the meanders is 5 to 14 mm, giving a ratio full course is commonly, though not exclusively, alternately of height/width between 2-5. The meanders are parallel to each winding and straight (modified from Ksiqzkiewicz, 1977). A tlantic G eology 231

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Fig. 5. Trace fossils from the Wapske Formation. (A) Helminthoida miocenica (TF.F111), x 1.64. (B) Helminthoida cf. labyrinthica (TF.F178), x 1.20. (C) Helminthopsis abeli (NBMG 9675), x 1.19. (D) Helminthopsis cf. granutata (NBMG 9682) (arrowed), x 0.91. (E) Helminthopsis isp. (NBMG 9683), x 0.51. (F) Helminthopsis hieroglyphica (NBMG 9680) (arrowed), x 1.1. All specimens are preserved in convex hyporelief. 232 H an and P ickerill

Description and Remarks: See Han and Pickerill, 1995. Remarks: The sets of multiple ridges in the present specimen suggest the trace was produced by the digging action of an or­ Helminthopsis isp. ganism with multidigited appendages. The ridges are well-pre­ Figure 5E served in only one set which, however, permits confident as­ signment to Monomorphichnus multilineatus. Material: One specimen (NBMG 9683) and 67 specimens col­ lected and over one hundred field occurrences. Ichnogenus Neonereites Seilacher, 1960

Description and Remarks: See Han and Pickerill, 1995. Type ichnospecies: Neonereites biserialis Seilacher, 1960

Ichnogenus Hormosiroidea Schaffer, 1928 Although Seilacher and Meischner (1965) considered Neonereites a preservational variant, though not strictly syn­ Type ichnospecies: HormosiroideaJlorentina Schaffer, 1928 onymous, with the associated ichnogenera Nereites MacLeay, Hormosiroidea cf. beskidensis (Plicka, 1974) Scalarituba Weller and Phyllodocites Geinitz, most recent au­ Figure 6A thors have preferred to retain it as a distinctive ichnotaxon (e.g., Benton, 1982b; Narbonne and Aitken, 1990; Fillion and Material: Three specimens (TF.F176, TF.F177, TF.F 209). Pickerill, 1990; Crimes and Crossley, 1991). We follow the opin­ ion of these latter authors, more complete reasoning for which Description: Specimens are preserved in convex hyporelief on is given in Pickerill (1991). the sole of a ripple-marked, 30 mm thick, buff, parallel-lami­ nated siltstone. They consist of two parallel rows of circular or Neonereites biserialis Seilacher, 1960 elongate knobs, each 2 to 7 mm in diameter. Individual rows 10 Figure 6E to 19 mm apart; the interval between successive individual knobs, 3 to 8 mm, is not constant along the length. The knobs Material: Seven specimens (TF9202-13, TF9206-1-10, are arranged alternately, in pairs, or irregularly. TF9208-11, TF9209-10, TF9210-11, TF9210-2-10).

Remarks: The nomenclature of Hormosiroidea (and the mor­ Description: Straight to winding biserial rows of circular to phologically similar ichnotaxon Saerichnites Billings) has been slightly ovate pustules preserved in convex hyporelief on the discussed by Seilacher (1977) and Crimes and Anderson (1985) soles of 23 to 60 mm thick, grey siltstones or fine-grained sand­ and there is little need for further discussion. It is a form that stones. The two rows are closely spaced or slightly separated by occurs both in deep-water (Seilacher, 1977; Crimes etal., 1981; an axial groove. Individual traces are up to 115 mm long and Crimes and Crossley, 1991) and shallow-water (Frey and 10 to 22 mm wide. Though typically biserial some specimens Chowns, 1972; Crimes and Anderson, 1985; Walter etal., 1989) may have only one or three pustules arranged in places. The marine environments, though mostly in flysch deposits. individual pustules are 4 to 8 mm in diameter. Depending on the degree of erosion, Hormosiroidea can exhibit considerable morphological variation (Seilacher, 1977). Remarks. The predominant arrangement of the pustules The alternation of knobs in material from the Wapske Forma­ biserially permits assignment to N. biserialis. tion is not as well developed as in Plicka’s (1974) specimen of H. beskidensis. Additionally, in some parts of the specimens Neonereites multiserialis Pickerill and Harland, 1988 only a single row of burrow knobs is present. The present speci­ Figure 6C mens are therefore only tentatively identified as Hormosiroidea beskidensis. Material: Thirteen specimens (TF9201-10, TF920I-11, TF9208-10, TF9208-21, TF9209-11, TF9210-10, TF9210-12, Ichnogenus Monomorphichnus Crimes, 1970 TF9210-13, TF9210a-10, TF.F100 (partim)) and several occur­ rences in the field. Type ichnospecies: Monomorphichnus bilinearis Crimes, 1970 Description: Individual specimens comprise three, up to four, Monomorphichnus multilineatus Alpert, 1976 rows of interconnected pustules preserved in convex hyporelief Figure 6B on the soles of 12 to 58 mm thick, grey or brown, parallel- laminated siltstones or fine-grained sandstones. The burrows Material: One specimen (NBMG 9352 (partim)). are horizontal, unbranched and longitudinally curved to irregu­ larly sinuous. Individual rows are up to 100 mm long and 11 to Description: Trace preserved in convex hyporelief on the sole 24 mm wide. Pustules are smooth and unornamented, spheroi­ of a 13 mm thick, parallel-laminated, fine-grained sandstone dal to ellipsoidal, variable in size (3 to 6 mm in diameter). Indi­ as three to seven parallel, very low, straight ridges or dig marks, vidual pustules may occur discretely or be amalgamated. Verti­ 0.5 mm wide and 6 to 14 mm long with an interval of 0.4 mm cal transverse sections of the specimen show that the pustules between adjacent ridges, forming sets 2 to 3 mm deep and 4 to are hemispheres. Burrow-fill is massive and similar to the host 6 mm wide. The central ridges are deeper and of the same width rock. as the lateral ones. A tlantic G eology 233

Fig. 6. Trace fossils from the Wapske Formation. (A) Homiosiroidea cf. beskidensis (TF.F177), x 1.0. (B) Monomorphichnus multilineatus (NBMG 9352 {partim)), x 1.0. (C) Neonereites multiserialis (TF9201-11), x 1.11. (D) Neonereites uniserialis (TF.F100), x 1.23. (E) Neonereites biserialis (TF9206-1-10), x 1.0. (F) Palaeophycus heberti (TF.F225), x 0.84. (G) Palaeophycus strialus (arrowed) and Palaeophycus isp. (TF.F218), x 1.0. (H) Palaeophycus tubularis (TF.F129), x 1.0. All specimens are preserved in convex hyporelief. 234 H an and P ickerill

Remarks: Pickerill (1991) has discussed this ichnospecies in collapse and show central furrows and two narrow walls. Bur­ detail based on material previously collected from the Wapske rows overlap or run alongside each other to create false branch­ Formation. Specimens described herein, and more recently col­ ing. lected, support his conclusions. Remarks. The present specimens are classified as P. heberti Neonereites uniserialis Scilacher, 1960 based on their characteristic thickly lined, unornamented walls Figure 6D which distinguish this ichnospecies from P. tubularis. The lack of any external surface ornamentation distinguishes P. heberti Material: Three specimens (TF112, TF.F100 (partim), TF.F103) from all other ichnospecies of Palaeophycus (Pemberton and and several occurrences in the field. Frey, 1982; Frey and Pemberton, 1991).

Description: Curved to irregularly winding, uniserial rows of Palaeophycus striatus Hall, 1852 circular to subcircular pustules preserved in convex hyporelief Figure 6G on the soles of 25 to 30 mm thick, massive or parallel-lami­ nated, siltstones and fine-grained sandstones. Individual rows, Material: Six specimens (TF9208-1-12, TF9209-19, TF9210- up to 730 mm long, are composed of pustules of 2.5 to 3.5 mm 4-10, TF129, TF.F206, TF.F218 (partim)) and several field oc­ diameter typically identical throughout their exposed length. currences. Pustules, however, may be connected with Helminthopsis isp. to form a compound specimen. Individual pustules are smooth Description: Horizontal to slightly inclined unbranched bur­ and closely connected or occasionally mutually overlap. rows, straight or slightly sinuous, covered with faint, parallel, longitudinal striae, preserved on the soles and in full relief of Remarks: Neonereites uniserialis diifers from N. biserialis and 35 to 50 mm thick, greenish grey, grey, buff, parallel-laminated N. multiserialis in being uniserial and from Microspherichnus siltstones or fine-grained sandstones. Burrow-fill is massive and linearis Hakes in being preserved in convex hyporelief. The similar to the host rock. Longitudinal striae consist of multiple compound (sensu Pickerill, 1994) specimen has not been previ­ and yet continuously aligned minute circular individual dots. ously described in the literature. Its occurrence possibly sug­ Transverse cross-sections show that the burrows are circular. gests that the individual tracemaker could produce different Burrows are 5 to 19 mm wide and up to 150 mm long. Burrow structures reflecting different behavioural patterns (cf. Bromley, width is generally constant along the length but may swell or 1990). narrow in places, especially where entering an interface.

Ichnogenus Palaeophycus Hall, 1847 Remarks: The massive burrow-fill suggests that it has not been sorted or processed by the tracemaker. The continuous and lon­ Type ichnospecies: Palaeophycus tubularis Hall, 1847 gitudinal parallel striae distinguish P. striatus from other stri­ ate ichnospecies of Palaeophycus (Pemberton and Frey, 1982). The nomenclatural history of Palaeophycus and the mor­ phologically similar ichnotaxon Planolites Nicholson has been Palaeophycus tubularis Hall, 1847 recently reviewed by Keighley and Pickerill (1995b). These au­ Figure 6H thors recommended that despite the inherent difficulties, the nomenclatural scheme comprehensively proposed by Pemberton Material: Fourteen specimens (TF9201-14, TF9201-19, and Frey (1982) is the most workable and, therefore, is that TF9201-20, TF9202-14, TF9209-19, TF129, TF.F126-TF.F129, adopted herein (see also Fillion, 1989; Fillion and Pickerill, TF.F164—TF.F167) and many more occurrences in the field. 1990). Description: The burrows are preserved in convex hyporelief Palaeophycus heberti (de Saporta, 1872) on the soles of, or in endorelief within, 25 to 50 mm thick, buff, Figure 6F brown, grey, parallel- or cross-laminated siltstones and fine­ grained sandstones. The burrows are unbranched, straight to Material: Four specimens (TF.F161, TF.F224 (partim), TF.F225 slightly curved, horizontal to undulatory, cylindrical, elliptical (partim), TF.F226 (partim)) and several occurrences in the field. or flattened, 4 to 12 mm in diameter and up to 120 mm long. Wall-lining, where preserved, is thin; burrow-fill is similar to Description: Straight to slightly curved, cylindrical to flattened, host rock and typically massive. Burrow surfaces are typically horizontal, unbranched, smooth-walled, lined burrows, 4 to 11 smooth. mm wide and up to 105 mm long, preserved in convex hyporelief on 27 mm thick, greenish grey, buff, cross- to parallel-lami­ Remarks: These morphologically simple burrows are assigned nated siltstones and fine-grained sandstones. The well-devel­ to P tubularis despite the fact that the burrow linings are diffi­ oped wall-lining is 1 to 2 mm thick and darker in colour than cult to detect. Thin wall-linings are easily removed by weather­ the burrow-fill which is massive and similar to the host rock. ing, commonly resulting in seemingly unlined burrows, which, Burrow width remains constant though some burrows exhibit nevertheless, should be assigned to this ichnospecies. A tlantic G eology 235

Palaeophycus isp. Ichnogenus Phycodes Richter, 1850 Figure 6G Type ichnospecies: Phycodes circinatus Richter, 1853 Material: Three specimens (TF124, TF9208-42, TF.F218 Phycodesflabellus M iller and Dyer, 1878 ipartim)) and numerous specimens collected and observed in Figure 7E the field. Material: Three specimens (TF9207-10, TF.F185, TF.F186). Description: Cylindrical to ellipsoidal, seemingly unlined or thinly lined, horizontal to inclined burrows, preserved in con­ Description: The burrow systems are preserved in convex vex hyporelief on the soles of, or endorelief within, siltstones hyporelief on the soles of 18 to 28 mm thick, grey, parallel- and fine- to medium-grained sandstones. The straight to slightly laminated siltstones and fine-grained sandstones. The speci­ curved burrows are 1 to 8 mm in width and variable in length. mens are 30 to 40 mm long, 10 to 25 mm wide and 2 to 5 mm Burrow-fill is massive and similar to the host rock. deep. The proximal branches are initially parallel and then project out at very acute angles in a flabellate or broomlike fash­ Remarks: Poor and incomplete preservation, as well as lack of ion. The entire systems are composed of 3 to 7 branches which knowledge of the three-dimensional character of the specimens, are 2 to 5 mm wide. The branches are close to each other, straight precludes ichnospecific assignment. to slightly curved. The burrows are faintly annulated and some portions are smooth or covered with small dots. The proximal Ichnogenus Paleodictyon Mcneghini in Murchison, 1850 portion of the burrows consists of a few tunnels. Burrow-fill is similar to the host rock and massive. Type ichnospecies: Paleodictyon strozzi Meneghini in Murchison, 1850 Remarks: Osgood (1970) discussed this ichnospecies in detail. Paleodictyon isp. It is difficult to generalize on an absolutely distinctive pattern Figure 7A for Phycodes flabellus, because the ichnospecies exhibits much variation in overall shape. P. flabellus differs from the morpho­ Material: Three specimens (TF9208-1-14, TF9208-1-16, logically similar P circinatus Richter by its lack of a spreite, TF9208-1-17). and the presence of a more shallow, flabellate burrow network.

Description: String-sized networks preserved in convex Phycodes palmatus (Hall, 1852) hyporelief on the soles of 26 to 60 mm thick, grey, parallel- Figure 7H laminated, fine-grained sandstones. The poorly preserved speci­ mens consist of irregular or regular polygons. Though no com­ Material: Five specimens (TF9208-21, TF9208-22, TF9210- plete horizontal meshes are present, hexagonal meshes can be 13, TF9306-5, TF128). inferred. Individual polygons are, diagonally, approximately 25 to 45 mm in diameter and of the same shape. Strings are cylin­ Description: The burrow systems are preserved as horizontal drical to subcylindrical and vary in thickness from 2 to 3 mm palmate structures, 40 to 55 mm in overall width, on the soles depending on height. Strings are characteristically straight and of 28 to 60 mm thick, grey, parallel-laminated siltstones and smooth; course change in strings is typically sharp and no ver­ fine-grained sandstones or interlayered sandstones and mud­ tical outlets are observed. stones. Each consists of a smooth proximal burrow, from which project, at acute angles, two to seven burrows, each 7 to 10 mm Remarks: Classification of Paleodictyon at the ichnospecific in diameter and 20 to 55 mm long. The branches, straight or level is still plagued by different authors adopting different cri­ very slightly curved, are close to or depart slightly from each teria (McCann and Pickerill, 1988; Pickerill, 1990; Crimes and other. Burrow surfaces may be covered by minute tubercles. Crossley, 1991). Mesh size, regularity, and thickness of string diameter (Ksiazkiewicz, 1970, 1977) or only mesh shape Remarks. Specimens described here closely resemble Hall’s (Seilacher, 1977) have been considered important in the dis­ (1852) syntypes and, accordingly, are assigned to P palmatus. tinction of ichnospecies. The main problem with each of these Although a spreite is not present, the palmate branching pat­ schemes is the complete gradation between different net sizes, tern is clearly obvious. string diameters and, indeed, net shapes (Crimes and Crossley, 1991). Therefore, a taxonomic revision of the ichnogenus is Phycodes aff. palmatus (Hall, 1852) still warranted (McCann and Pickerill, 1988; Pickerill, 1990; Figure 7B Crimes and Crossley, 1991). The generally incomplete preser­ vation of the material described herein precludes ichnospecific Material: Seven specimens (TF9209-16, TF9209-22, TF9209- assignment. 24, TF9209-25, TF.F187, TF.F204).

A tlantic G eology 237

Description: Horizontal burrow systems preserved in convex ing fashion is similar to that found in the ichnogenus Treptichnus hyporelief on the soles of 11 to 45 mm thick, grey, argillaceous Miller which is probably closely related to Phycodes pedum siltstones and fine-grained sandstones. The overall pattern of (Banks, 1970). the burrows is thicket-shaped. The systems, 40 to 100 mm wide and 40 to 80 mm long, consist of 5 to 8 individual cylindrical Phycodes templus Han and Pickerill, 1994 branches that project outward from a centre. The proximal por­ Figure 7C tions are not well-preserved or even absent. The branches, smooth, straight or slightly or strongly curved, are 35 to 65 mm Material: Eight specimens (NBMG 9202 - NBMG 9209) and in length and 5.5 to 7 mm in width which is constant or slightly other collected specimens. tapers distally. The elongate branches are closely spaced proxi- mally and widely spaced distally, between 1 and 22 mm apart Diagnosis: Phycodes possessing two or, more typically, several (mostly 10 mm apart). horizontally interconnected broomlike or flabellate bundles that collectively form an inverted pagoda-like structure. Remarks. The present specimens differ from typical P. palmatus by individual branches being more widely spaced, elongate, Description and Remarks: See Han and Pickerill, 1994a. strongly curved and lacking a spreite. However, the overall pat­ tern is similar to P. palmatus; therefore, the specimens are as­ Ichnogenus Protopaleodictyon Ksi^ikiewicz, 1970 signed to P. aff. palmatus. Type ichnospecies: Protopaleodictyon incompositum Phycodespedum Scilacher, 1955 Ksiazkicwicz, 1970 Figure 7D Protopaleodictyon incompositum Ksi^zkiewicz, 1970 Figure 7F Material: Three specimens (TF9210-14, TF102, TF.F191). Material: One specimen (TF9208-1-15). Description: Burrow systems preserved in convex hyporelief or full relief on the soles of 19 to 28 mm thick, grey siltstones Description: Burrow system preserved in convex hyporelief on and fine-grained sandstones. The burrows consist of a straight the sole of a 60 mm thick, grey, parallel-laminated, fine-grained row of short, corn-like branches which are of similar size and sandstone. The burrow system consists of wide first-order me­ arranged in a zigzag fashion or closely spaced and bundled in a anders and sinuous second-order meanders with apical append­ fan-shape, with branches projecting in a similar direction. No ages. The first-order meanders are only partially preserved. The central stem is present. The cylindrical to subcylindrical appendages are of variable length, 3 to 15 mm, and are posi­ branches alternately emerge or branch out from the inner side tioned on the apices of the second-order meanders and never of previous branches at angle of 50° to 70° and form a zigzag form a mesh; only one appendix branches from the apex on the shape. The surfaces of the branches are transversely ringed with convex side of the meanders. The second-order meander have a one rounded to subrounded shallow, narrow furrow which cuts wavelength of 7 to 16 mm and amplitude of 2 to 3 mm. The the branches into two equal-length parts. The branches, 9 to 11 strings, 1.5 to 2 mm wide, are smooth. mm long and 5 to 6 mm wide at midlength, taper gradually outward and form sharp terminations. The distal parts of the Remarks: Similar to Paleodictyon, there is still no general branches are less deeply impressed than the proximal parts. The agreement as to which characters constitute a distinctive incomplete specimens have lengths of 46 to 65 mm and widths ichnospecies of Protopaleodictyon (Pickerill, 1981; Pickerill et of 8 to 16 mm. Burrow-fill is the same as the host rock. No al., 1982). Seilacher (1977) utilized the number of branches/ sickle-like gallery is observed. Cross-cutting reveals that the undulations and the presence of first- and second-order mean­ individual branches are almost on the same plane and mud- ders, while Ksiqzkiewicz (1977) employed the regularity and lined. spacing of the first-order meanders, and the size and thickness of the strings. Ksiqikiewicz’s (1977) scheme is that adopted R em arks: Phycodes pedum was originally described by herein in view of the obvious variability exhibited in the speci­ Seilacher (1955) as a semicircular form; however, meandering mens figured by him. The ichnogenus has been ascribed to a and more or less straight and looping forms have each been burrowing deposit feeder, possibly an infaunal annelid (Tanaka, subsequently described (Crimes et al., 1977; Bryant and 1971; Kern, 1980; Pickerill, 1981). Pickerill, 1990). The minor branches normally all occur on the The specimen from the Wapske Formation rarely shows same side of the main burrow but specimens with one or more first-order meanders, but the second-order meanders, typically on the opposite side to the remainder (Crimes et al., 1977) and with apical appendages, differentiates it from Cosmorhaphe. P. even alternating dextral and sinistral branches (Biyant and incompositum is distinguished from the morphologically simi­ Pickerill, 1990) have been reported. In the latter case, the branch­ lar ichnospecies P. minutum Ksiqzkiewicz by its larger size and

Fig. 7. Trace fossils from the Wapske Formation. (A) Paleodictyon isp. (TF9208-1-17), x 0.68. (B) Phycodes aff. palmatus (TF9209-22), x 1.22. (C) Phycodes templus (NBMG 9202), x 0.60. (D) Phycodes pedum (TF102), x 1.06. (E) Phycodesjlabellus (TF.F186), x 2.28. (F) Protopaleodictyon incompositum (TF9208-1-15), x 1.0. (G) Protovirgidaria dichotoma (NBMG9348 (right one) and NBMG 9349 (left one), x 0.82. (H) Phycodes palmatus (TF9208-21), x 1.22. All specimens are preserved in convex hyporelief. 238 H an and P ickerill from P. submontanum (Azpeitia Moros) by its more regular me­ parallel-laminated siltstones or fine-grained sandstones. Bur­ ander pattern of the first-order meanders and also by the latter’s row walls are either sharp and smooth or indistinct. The bur­ much longer appendages and a more pronounced tendency to­ row-fill is massive and may be finer or coarser in grain size and ward formation of networks. darker or lighter in colour than the surrounding matrix.

Ichnogenus Protovirgularia M ’Coy, 1850 Remarks: Although Alpert’s (1974, 1975) nomenclatural scheme for the main ichnospecies of Skolithos is still regarded Type ichnospecies: Protovirgularia dichotoma M ’Coy, 1850 as the most satisfactory (Fillion and Pickerill, 1990), its tax­ Protovirgularia dichotoma M ’Coy, 1850 onomy is still in need of revision. With respect to this scheme, Figure 7G specimens from the Wapske Formation are identical to S. linearis and are identified accordingly. Material: Nineteen specimens (NBMG 9334 - NBMG 9352). Ichnogenus Taenidium Heer, 1877 Emended diagnosis: Unbranched, keel-like trail, typically, but not universally, with a median ridge or furrow from where Type ichnospecies: Taenidium serpentinum Heer, 1877 paired, lateral, wedge-shaped appendages, commonly only a few Taenidium serpentinum Heer, 1877 millimetres in length, and of even or variable spacing, origi­ Figure 8B nate. Lateral appendages may be normal or at an acute angle to the median ridge or furrow (modified after Volk, 1961; Benton, Material: Two specimens (TF.F206, TF.F235). 1982a). Description: Unlined, or very thinly mudstone lined, cylindri­ Description and Remarks. See Han and Pickerill, 1994b. cal serpentiform burrows, possessing a well-spaced arcuate meniscate backfill and preserved in convex hyporelief on the Ichnogenus Rusophycus Hall, 1852 soles of 45 and 75 mm thick, grey, brown, parallel-laminated, fine-grained sandstones. The burrows are straight or slightly Type ichnospecies: Rusophycus clavatus Hall, 1852 curved, unbranched, 4 mm wide and 100 mm long. Termina­ Rusophycus cf. carbonarius Dawson, 1864 tions are obscure. The outer surfaces are wavy or slightly Figure 8A annulated. The faint annulations appear to correspond to the menisci of the fills. In places where the grey mudstone has been Material: One specimen (TF9208-1-34). removed by weathering, the inner surfaces are obviously annulated by menisci which are 4 mm wide and 3.5 to 4 mm Description: Small, bilobate bean-like lobes preserved in con­ long. vex hyporelief on the sole of a 55 mm thick, grey, parallel-lami­ nated, fine-grained sandstone. The lobes taper very slightly an­ Remarks: The ichnotaxonomy of simple, unbranched meniscate teriorly. Maximum width is 10 mm, near the mid-length, and burrows has been reviewed by D ’Alessandro and Bromley (1987) tapers to only 2.5 mm. Maximum length is 17 mm, giving a and more recently by Keighley and Pickerill (1994). With re­ length/width ratio 1.7. The median longitudinal furrow is shal­ spect to these revisions the material described herein can confi­ low and narrow, being deepest near the mid-length of the lobes. dently be assigned to T. serpentinum. T. cameronensis (Brady) Surface of the lobes possess very poorly preserved transverse differs by possessing intermeniscate segments distinctly longer scratch marks, each 1 mm in width. than wide, T. satanassi D ’Alessandro and Bromley possesses only weakly arcuate menisci and an obviously pelleted fill (see Remarks: The ichnotaxonomy of small cruzianids and D’Alessandro and Bromley, 1987) and T. barretti (Bradshaw) rusophycids has recently been reviewed by Keighley and Pickerill possesses deeply arcuate or hemispherical, tightly packed me­ (1995a). Although poorly preserved, the specimen described nisci. herein closely resembles R. carbonarius and is tentatively iden­ tified as such. Ichnogenus Uchirites Macsotay, 1967

Ichnogenus Skolithos Haldcman, 1840 Type ichnospecies: Uchirites triangularis Macsotay, 1967 Uchirites implexus Rindsberg, 1994 Type ichnospecies: Fucoides? linearis Haldcman, 1840 Figure 8D Skolithos linearis Haldcman, 1840 Figure 8C Material: Seven specimens (TF9201-15, TF9201-16, TF9207- 11, TF9208-24, TF.F182, TF.F183). Material: Four specimens (TF9210-4-11, TF9302-01, TF122, TF.F203). Description: Straight, curved or meandering, triangular, hori­ zontal structures, preserved in convex hyporelief on the soles of Description: Straight, vertical to slightly inclined, unbranched, 24 to 45 mm thick, grey, brown or orange, parallel- and cross- cylindrical burrows, 3 to 7.5 mm in diameter and of uncertain laminated siltstones and fine-grained sandstones. Specimens length, preserved in endorelief within 20 to 40 mm thick, grey, possess a single median crest on the lower (ventral) surface and A tlantic G eology 239

Fig. 8. Trace fossils from the Wapske Formation. (A) Riisophycus cf. carbonarius (TF9208-1-34), x 2.0. (B) Taenidium serpentinum (TF.F206) (arrowed), x 1.63. (C) Skolithos linearis (TF9302-01), x 1.13. (D) Uchirites implexus (TF.F183), x 0.46. (E) Umfolozia cf. sinuosa (TF.F235), x 1.61. (F) Track form B (TF9203-11), x 1.11. (G) Scratch marks (TF.F194), x 1.11. (H) Track form A (TF.F210), x 0.97. C is preserved in endorelief; E in concave relief; remainder in convex hyporelief. 240 H an and P ickerill paired, lateral, smooth marginal flanks. The crests, 1.5 to 2 Description: Trackway preserved in convex hyporelief on the mm wide, are typically axial but may be slightly disposed to surface of a greenish grey siltstone. The straight trackway con­ one side probably as a result of compaction. Structures are 57 to sists of two parallel rows or series of circular impressions. It is 320 mm in length and 12 to 15 mm in width which is constant 120 mm long and 5 to 6 mm wide, individual impressions be­ in single examples. In two specimens, terminations are com­ ing 1 mm in diameter and usually 5 to 6 mm apart in successive posed of oval-shaped bulges that lack both the median crest and rows, rarely 4 mm apart. The impressions occur commonly in the inclined lateral lobes but are wider and higher than the main pairs, more rarely dispersed. The area between the two rows is portion of the structures. In one specimen, the median crest undisturbed. terminates 10 mm before the bulging termination. The fill is similar to or slightly coarser than the surrounding host rock but Remarks: Morphologically, this trackway is similar to examples does not disturb the horizontal laminae in and above the un­ of Honnosiroidea beskidensis. However, the latter is much larger lined structures. in size and possesses vertical outlets arranged alternately. The trackway was probably produced by an arthropod-like, bilater­ Remarks: Rindsberg (1994) has recently reviewed the ally symmetrical animal. ichnotaxonomy of Uchirites and, in so doing, formulated the ichnospecies U. implexus. The present material differs only from Track form B his types in the absence of delicate lateral striations, though we Figure 8F regard the absence of these in the Wapske Formation material as preservational. Otherwise, the material is identical and is Material: One specimen (TF9203-11). therefore considered conspecific. Description: Preserved in convex hyporelief on the sole of a 32 Ichnogenus Umfolozia Savage, 1971 mm thick, grey, fine-grained sandstone. The trace is composed of two contrasting morphologies, an axial feather-like form and Type ichnospecies: Umfolozia sinuosa Savage, 1971 two lateral parallel rows of paired foot or claw imprints appear­ Umfolozia cf. sinuosa Savage, 1971 ing as small circular knobs. The whole trace is 62 mm long and Figure 8E 28 mm wide. The slightly curved axial feather-like trail consists of a Material: Two specimens (TF.F235, TF.F237). ‘stem’ and lateral fine ‘filaments’. The ‘stem’ is 0.5 to 1.0 mm wide. The fine ‘filaments’ propagate from the ‘stem’ and form Description: Simple trackways preserved in concave relief, re­ a V-shape with angles of 16° to 22° and individual arms are 0.5 spectively, on the surfaces of 29 and 80 mm thick, grey, paral­ mm wide and 8 to 10 mm long. The ‘stem’ may be formed by lel-laminated, fine-grained sandstone and siltstone. The overlapping ‘filaments’. The axial trail is 4 to 5 mm wide. The trackways, 55 to 90 mm long and 8 to 12 mm wide, consist of ‘filaments’ have no branches. two, more or less parallel, straight rows of imprints. Individual The two rows of foot or claw imprints on each side of the imprints, 0.5 to 1 mm wide and up to 4 mm long, are arranged axial trail are roughly parallel and symmetrical. The foot im­ approximately in cycles of five and are circular or tadpole­ prints are clear and distinct, circular, large or small, unifid or shaped, having a circular deep impression and an elongate shal­ bifid, and 2 to 4 mm in diameter with intervals of 4 to 9 mm, up low or deep tail. The paired imprints are 4 to 9 mm apart and to 11 mm, in a successive row. The distance between rows of successive imprints are separated by a distance of 3 to 9 mm. imprints and the axial trail is between 5 and 12 mm. The cen­ Individual imprints are transverse or oblique to the axial line, tral trail is slightly curved while the lateral rows of imprints oriented at angles of 40° to 50°. The tails in both rows of remain virtually straight. trackways are oriented toward the same direction. Remarks: The axial part of this trackway is similar to both Remarks: The significant characteristic of Umfolozia, a ‘Chloephycus' Miller and Dyer (Osgood, 1970, pi. 80, figs. 1, trackway suggested by Savage (1971) and Anderson (1981) to 6) and inorganic groove moulds and chevron marks (D2ulyAski have been produced by peracarid crustaceans, is the repetitive and Walton, 1965, p. 88, fig. 61; p. 100, figs. 68B, C). The nature of the tracks comprising five pairs of appendages and present specimen, however, is believed to be biogenic in origin the presence or otherwise of a sinuous series of small oval tel- based on the following observations: son marks (Anderson, 1981). The present samples have a diag­ (1) The central feather-like marks are curved, unlike typi­ nostic track cycle of five, though this is not completely devel­ cal ‘Chloephycus’ or inorganic groove markings; oped along their entire length. Additionally, there are no longi­ (2) On the same slab, another poorly preserved specimen tudinal drag lines or drag marks. Nevertheless, we tentatively shows similar feather-like features in which the V-angles point regard the material as U. sinuosa. in an opposite direction to the described specimen; (3) The feather-like marks are flanked by two rows of tracks Track form A which are more or less parallel and possess scratches; Figure 8H (4) Other scratch marks occur on the same stratification surface. Material: One specimen (TF.F210). Given a biogenic origin, this trackway does invite com­ A tlantic Geology 241

parison with several previously described ichnotaxa. For ex­ ------1976. Trilobite and star-like trace fossils from the White-Inyo ample, Protovirgularia M’Coy is an ichnotaxon that only has Mountains, California. Journal of Paleontology, 50, pp. 226-239. the central feather-like parts. Protichnites Owen has two rows Anderson, A.N. 1981. The Umfolozia arthropod trackways in the Per­ of bifid or trifid imprints which flank the median double straight mian Dwyka and Ecca Series of South Africa. Journal of Paleon­ tology, 55, pp. 84-108. trails and Mesichnium Gilmore has two parallel rows of foot­ Azpeitia M oros, F. 1933. Datos para es estudio paleontologico del prints with a median row of suboval regularly spaced depres­ Flysch de la Costa Cantabrica y de algunos otros puntos de Espaiia. sions. Boletin del Instituto Geologico y Minero de Espaiia, 53, pp. 1-65. Imprints of this type were probably made by an unknown B andel, K. 1973. Trace fossils from the Upper Devonian Nehden Silt- bilaterally symmetrical arthropod which had vertical append­ stone of Wuppertal-Barmen (Nordrhein-Westfalen, Germany). ages resulting in the central feather-like drag marks, and paired Palaeontographica, Abteilung A, Palaozoologie-Stratigraphie, 142, feet leaving lateral rows of scratch marks. Assignment to previ­ pp. 156-176. ously described ichnotaxa is, however, considered unwise pend­ B anks, N.L. 1970. Trace fossils from the late Precambrian and Lower ing discovery of additional or more informative material. Cambrian of Finnmark, Norway. In Trace fossils. Edited by T.P. Crimes and J.C. Harper. Geological Journal, Special Issue 3, Seel House Press, Liverpool, pp. 19-35. Scratch marks B enton, M.J. 1982a. Trace fossils from Lower Palaeozoic ocean-floor Figure 8G sediments of the Southern Uplands of Scotland. Royal Society of Edinburgh Transactions: Earth Sciences, 73, pp. 67-87. Material: One specimen (TF.F194). ------1982b. Dictyodora and associated trace fossils from the Palaeozoic of Thuringia. Lethaia, 15, pp. 115-132. Description: A series of imprints are preserved in convex B illings, E. 1872. On some fossils from the Primordial rocks of New­ hyporelief on the surface of a 33 mm thick, grey, fine-grained foundland. Canadian Naturalist and Quarterly Journal of Science sandstone. The imprints consist of rectangular impressions, in­ with the Proceedings of the Natural History Society of Montreal, New Series, 6, pp. 465-479. dividually 3 mm wide and 5 to 7 mm long, with a shallow, B oucot, A.J. and W ilson, R.A. 1994. Origin and early radiation of short furrow on their lower portions. Individual impressions terebratuloid brachiopods: thoughts provoked by Prorensselaeria are parallel to each other but all align at 30° to 35° with the and Nanothyris. Journal of Paleontology, 68, pp. 1002-1025. presumed axis of movement. The specimen is 96 mm long and B radley, J. 1981. Radionereiles, Chondrites and Phycodesr, trace fos­ 5 to 6 mm wide, having intervals between successive imprints sils of anthoptiloid sea pens. Pacific Geology, 15, pp. 1-16. of 5 to 10 mm. B radshaw, M.A. 1981. Paleoenvironmental interpretations and sys- tematics of Devonian trace fossils from the Taylor Group (lower Remarks: MikulaS (1992) established the ichnogenus Beacon Supergroup), Antarctica. New Zealand Journal of Geol­ Interruptida for similar surface traces interrupted in regular in­ ogy and Geophysics, 24, pp. 615-652. B romley, R.G. 1990. Trace Fossils: Biology and Taphonomy. Unwin tervals and, indeed, the description, dimensions and illustra­ Hyman, London, 280 p. tion of Interruptida indet. B (Mikulas, 1992, pi. 13, fig. 5) are B rom ley, R.G. and A sgaard, U. 1979. Triassic freshwater similar to the present specimen. However, without more mate­ ichnocoenoses from Carlsberg Fjord, east Greenland. rial we are reluctant to accept MikulaS’s (ibid.) taxonomic de­ Palaeogeography, Palaeoclimatology, Palaeoecology, 28, pp. 39- cision, as such examples could conceivably represent taphonomic 80. variants of previously established ichnotaxa. As such we retain B romley, R.G. and E kdale, A.A. 1984. Chondrites: a trace fossil indi­ the specimen in open nomenclature. cator of anoxia in sediments. Science, 224, pp. 872-874. ------1986. Composite ichnofabrics and tiering of burrows. Geo­ logical Magazine, 123, pp. 59-65. A cknowledgements B romley, R.G., P emberton, S.G., and R ahmani, R.A. 1984. A Creta­ ceous woodground: the Teredoliles ichnofacies. Journal of Pale­ We wish to thank A. Gomez, R. McCulloch and D. ontology, 58, pp. 488-498. Campbell for technical assistance, R. Wilson and L. Fyffe, New B rongniart, A.T. 1828. Histoire des vegetaux fossiles ou recherches Brunswick Department of Natural Resources and Energy, for botaniques et geologiques sur les vegetaux renfermes dans les assistance in defraying fieldwork expenses, and R. Miller, diverses couches du globe, volume 1. G. Dufour and E. d’Ocagne, Steinhammer Palaeontology Laboratory, New Brunswick Mu­ Paris, 136 p. seum, for providing appropriate repository numbers for several B ryant, I.D. and P ickerill, R.K. 1990. Lower Cambrian trace fossils from the Buen Formation of central North Greenland: prelimi­ specimens described and/or figured herein. Constructive cri­ nary observations. Gronlands geologiske Undersagelse Rapport, tiques of the initial manuscript were provided by H. Hofmann 147, pp. 44-62. and an anonymous reviewer. Financial support for this research C hamberlain, C.K. 1971. Morphology and ethology of trace fossils was provided by a Natural Sciences and Engineering Council from the Ouachita Mountains, southeast Oklahoma. Journal of of Canada operating grant to RKP, which is gratefully acknowl­ Paleontology, 45, pp. 212-246. edged. ------1977. Ordovician and Devonian trace fossils from Nevada. Bulletin of Nevada Bureau of Mines and Geology, 90, pp. 1-24. A lpert, S.P. 1974. Systematic review of the genus Skolithos. Journal C orbo, S. 1979. Vertical distribution of trace fossils in a turbidite se­ of Paleontology, 48, pp. 661-669. quence, Upper Devonian, New York State. Palaeogeography, ------1975. Planolites and Skolithos from the Upper Precambrian- Palaeoclimatology, Palaeoecology, 28, pp. 81-101. Lower Cambrian White-Inyo Mountains, California. Journal of C rimes, T.P. 1970a. The significance of trace fossils in sedimentology, Paleontology, 49, pp. 508-521. stratigraphy and palaeoecology, with examples from Lower 242 H an and P ickerill

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C. van Benthuysen, Albany, 362 p. organism traces on and in pelagic carbonates of the western equa­ H an, Y. 1995. Sedimentology and ichnology of the Lower Devonian torial Pacific. Palaeogeography, Palaeoclimatology, Palaeoecology, Wapske Formation, northwestern New Brunswick, eastern 23, pp. 263-278. Canada. Unpublished M.Sc. thesis. University of New Brunswick, E lliott, R.E. 1985. An interpretation of the trace fossil Cochlichnus Fredericton, New Brunswick, 372 p. kochi (Ludwig) from the East Pennine Coalfield of Britain. Pro­ H an, Y. and P ickerill, R.K. 1994a. Phycodes templus isp. nov. from ceedings of the Yorkshire Geological Society, 45, pp. 183-187. the Lower Devonian of northwestern New Brunswick, eastern Farres, F. 1967. Los “Dendrotichnium” de Espaiia. Institute Geologico Canada. Atlantic Geology, 30, pp. 37-46. y Minero de Espaiia. Notas y Comunicaciones, Boletin, 94, pp. ------1994b. 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